Seasonal maturity development of Baltic cod in different spawning areas: importance of the Arkona Sea for the summer spawning stock

We investigated the seasonal maturity development of cod in four areas of the Baltic Sea. Two different spawning peaks were identified and found to be consistent over the period 1992–2005. In the Kiel Bight and Mecklenburg Bight (ICES SD 22) a spawning peak was observed from March to April (spring spawning). In the areas of the Arkona Sea (ICES SD 24) and Bornholm Sea (ICES SD 25) the spawning peak occurred during summer. In the Bornholm Sea, the main spawning activities began in June and ended in September, with a spawning peak in June–August (summer spawning). In the Arkona Sea, which is a transition area between the Mecklenburg Bight and the central Baltic Sea, spawning began in March and lasted until July, with a spawning peak in June–July (summer spawning). Seasonal maturity development and proportions of spawning cod in June in the Arkona Sea were similar to that of the Bornholm Sea. In addition, the proportion of spawning cod in the Arkona Sea was positively correlated with the size of the spawning stock in the Bornholm Sea. Our results provide evidence of a spatial expansion of spawning activities of the summer spawning stock from the eastern Baltic Sea into the Arkona Sea. Therefore, the Arkona Sea should be considered as one of the spawning habitats of the summer spawning stock of Baltic cod.     

First evidence of spawning of eastern Baltic cod (Gadus morhua callarias) in the Belt Sea,the main spawning area of western Baltic cod (Gadus morhua L.)

Two cod stocks (western Baltic cod, WBC, and eastern Baltic cod, EBC) are managed in the Baltic Sea which is characterized by different main spawning areas and different main spawning periods. In this study we analyse the spatial and temporal occurrence of spawning individuals of both cod stocks in the main spawning grounds of the Baltic Sea based on eight microsatellite loci. Our results suggest that EBC (Gadus morhua callarias) has formed currently temporally stable, substantially homogeneous population not only in the Bornholm Sea (ICES SD: 25) but also in the Arkona Sea (ICES SD: 24). The presented analyses proved that EBC (G. m. callarias) can temporarily also spawn in the Belt Sea.     

Return of twaite shad <Emphasis Type="Italic">Alosa fallax</Emphasis> (Lacépède, 1803) to the Southern Baltic Sea and the transitional area between the Baltic and North Seas

The status of twaite shad was investigated within the Southern Baltic Sea (ICES subdivision 22–27) and transitional area between the Baltic and North Seas (division IIIa). The following sources of data were analysed: (i) commercial catch statistics and relevant publications, (ii) records from ichthyological museum collections, (iii) records from recent commercial and recreational fisheries, (iv) research fisheries with trawls. A total of 476 records of twaite shad including more than 16 million individuals were obtained for the time between the years 1836 and 2005. About 72.9% of all records originated from commercial catch statistics and publications, whereas 18.9% were received from ichthyological collections. Research fisheries provided 6.3%, and 1.9% of the records were obtained from recent commercial and recreational fisheries. Most records of twaite shad were estimated for subdivisions 24 (45.2%) and 26 (35.5%). From 1836 to 1959, 29.6% of the records date from the period until 1899. 70.4% of the records of twaite shad originate from the twentieth century until 1959. The mean annual catch of twaite shad between 1891 and 1959 amounted to 86,674 kg within subdivisions 24–26 of the Southern Baltic Sea. Catch data show an approximately 20-year-cyclicity of maximum yields and minimum catches, respectively. The maximum annual yield of twaite shad in subdivisions 24–26 (474,700 kg) was registered in 1940, the minimum annual yield was estimated in 1958 (10 kg). In the 1950s, the annual catches of twaite shad declined sharply. Until 1960 twaite shad catches and records originated mainly from the Pommeranian Bay/Pommeranian Coast and adjacent waters including the Szczecin Lagoon (subdivision 24), the Bay of Gdańsk, Vistula Lagoon and Vistula Spit (subdivision 26) and from the Curonian Lagoon and Curonian Spit (subdivision 26). The highest catches of twaite shad originated from the area of Curonian Lagoon/Curonian Spit from 1941 to 1960. Seasonal catches of twaite shad showed maximum values from May to July. From 1960 to 1989, only four records of A. fallax were registered in the Southern Baltic Sea which originated from German coastal waters in subdivisions 22 and 24. A total of 107 records of twaite shad was obtained in subdivisions 20–27 from 1990 to 2005. Most of these recent records originate from a twaite-shad stock in subdivisions 24, 25 and 26. There are indications which suggest a separate stock of twaite shad in subdivisions 20 and 21. Disappearance and recovery of twaite shad stocks were probably caused by the following factors: construction of barriers in rivers with spawning sites of twaite shad; habitat destruction in those rivers as consequence of gravel extraction and reengineering scheme to improve navigation and for flood defence purposes, water pollution in the lagoons of the Southern Baltic and in their tributaries, commercial fishery in the Southern Baltic and climatic variation in the Baltic Sea basin south of the latitude of 60° N. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: S. Dufour, E. Prévost, E. Rochard & P. Williot Fish and diadromy in Europe (ecology, management, conservation)     

Differentiation of western and eastern Baltic Sea cod stocks (Gadus morhua) by means of stable isotope ratios in muscles and otoliths

Stable isotope analysis of Baltic Sea cod from the Bornholm Sea (ICES‐SD 25) and western Baltic (Belt Sea, ICES‐SD 22) revealed significant differences in the δ¹⁵N and δ¹³C values of the dorsal muscle, as well as in the δ¹⁸O values of otoliths. The method pledges to become especially appropriate to the Baltic due to the high variability in oxygen isotope ratios associated with its estuarine nature.     

Towards better Understanding of Successful Management of the Gulf of Riga Herring Stock

The Baltic herring is one of the key elements of the pelagic ecosystem of the Baltic Sea, being of the most important commercial species of the sea. The structure and dynamics of herring populations (stocks) have been assessed and managed on international level since 1970s. Since 1990, all local herring populations have been combined and assessed as one stock (Central Baltic Herring in Sub-divisions 25–29 and 32). However, the continuously decreasing trend in stock biomass throughout almost 30-years period of observations indicates the failure of the stock management implementation of the scientific advice. The separate assessments of different stocks and results of hydro-acoustic surveys of different sub-units (herring in Sub-divisions 25–27, Sub-divisions 28, 29 and 32 and the Gulf of Riga herring) have revealed rather different situation within combined assessment unit: e.g. fishing mortality of the Gulf of Riga herring has decreased while a sharp increase in mortality is observed in the herring stocks of the North-eastern Baltic (Sub-divisions 28, 29 and 32). At the same time stock abundance and biomass of the Gulf of Riga herring have increased to record high in early 2000s allowing also higher catches. It is concluded that only separate analytical assessment of local populations combined with regular acoustical surveys and following different protection measures have proved to be a successful way in managing the herring stocks. Joint assessment and management of several populations with different structure and dynamics as one combined stock do not allow revealing the real situation and trends in its parts.     

Spatial and temporal variations in parasite prevalence and infracommunity structure in herring (Clupea harengus L.) caught to the west of the British Isles and in the North and Baltic Seas: implications for fisheries science

Herring Clupea harengus L. viscera were examined for endoparasitic infections as part of a multidisciplinary stock identification project (WESTHER, EU Contract no. Q5RS-2002-01 056) which applied a range of stock discrimination techniques to the same individual fishes to obtain comparable results for multivariate analysis. Spawning and non-spawning adults, and juvenile herring were caught, over 3 years, by commercial and research vessels from numerous locations to the west of the UK and Ireland, along with control samples of spawning fish from the eastern Baltic Sea, and juveniles from sites in the eastern and western North Sea, and the north of Norway. The metacercariae of two renicolid digeneans (Cercaria pythionike and Cercaria doricha), one larval nematode (Anisakis simplex s.s.) and one larval cestode (Lacistorhynchus tenuis) were selected as tag species. Results were compared with those from herring collected between 1973 and 1982, which suggested remarkable stability in the parasite fauna of herring in the study area. These species were used to compare the parasite infracommunities of spawning herring. A significant variation in infracommunity structure was observed between different spawning grounds. These results suggest that the parasite fauna of herring are spatially variable but remain temporally stable in both the short and long term. Significant differences in prevalence and abundance of infections and comparisons of parasite infracommunity enabled the separation of putative herring stocks west of the British Isles. Distinctive patterns of parasite infection in two different spawning groups off the north coast of Scotland suggest that this area is occupied by two spawning populations, one recruiting from the west of Scotland, the other from outside this area, and most likely from the eastern North Sea. The distribution patterns of L. tenuis, C. doricha and C. pythionike suggest the potential for fish that spawn in three distinct International Council for the Exploration of the Seas (ICES) management units to be present in mixed aggregations found over the Malin Shelf, with significant implications for management in this area.     

Effects of varying natural mortality and selectivity on the assessment of eastern Baltic cod (Gadus morhua Linnaeus, 1758) stock

In 2014, the International Council for the Exploration of the Sea (ICES) was unexpectedly unable to provide an analytical assessment of eastern Baltic cod stock; factors such as data issues, assessment methodology, and the ecological situation of cod were indicated as the reasons for this failure. Some evidence suggests that the natural mortality (M) of cod could increase substantially in forthcoming years and that the selectivity could change. In this paper, age‐structured and stock‐production assessment models were applied to simulate the dynamics of cod stock; in the models, both constant and increasing natural mortalities were permitted. In the age‐structured model, the effects of selectivity related to the cod size on the cod assessment were also analysed. In addition, stock with characteristics similar to Baltic cod stock and increasing natural mortality was generated and assessed with the age‐structured model using both constant and increasing M. It was shown that models with increasing natural mortality of cod in recent years perform much better than models with constant natural mortality in terms of the distribution of residuals and retrospective patterns. The models with size‐dependent selectivity did not perform better than other standard assessments. The assessment of generated stock (where natural mortality was increasing) with constant natural mortality in the assessment model showed a poor distribution of residuals and strong retrospective patterns, similar to the ICES assessment with constant M. The conducted simulations strongly suggest that the main reason for the poor recent cod assessment is the increase in natural mortality, which is not considered in the assessment methodology.     

Comparative analysis of alternative statistical models for differentiation of herring stocks based on meristic characters

A basic stock assessment problem is the mixing and separation of herring populations in their specific areas. Within the transition zone between the North and the Baltic seas (Skagerrak, Kattegat, The Sound) a mixing of two herring populations ( Clupea harengus L.) temporarily takes place. One major component stems from the Baltic Sea (spring spawner), and the other from the North Sea (autumn spawner); each stock exhibits different meristic characteristics. In order to separate the two herring populations, 'pure' learning samples of meristic characters were created in 1995 as differential variables. Mainly gained during two research surveys at two geographically remote areas where a mixing of herring components was relatively unlikely, these learning samples were considered as representative for one or the other herring population. This paper compares the current (ICES) separation approach for herring stock assessment with two alternative methods wherein vertebra counts (vc) are used as meristic characters. The two proposed methods are (a) an inverted variance weighted linear model, and (b) a separation rule based on a quadratic discriminant analysis. The paper summarizes and discusses the results of predicting fractions of the two herring components derived from all three separation models. The underlying example data set stems from a 1991–97 routine Swedish survey in the transition zone. In comparison, the quadratic discriminant analysis separation model was identified as superior to the two other methods. Furthermore, the model suggests a higher degree of mixing of the two herring stocks in the transition area than was previously thought; inter-annual changes in the geographical distribution of the two populations are suggested as being less variable than previously assumed.     

The potential fecundity of cod in the Baltic Sea from 1993 to 1999

Investigations on the reproduction biology of cod in the Baltic Sea (Gadus morhua L. and G. morhua callarias L.) were carried out beginning in 1993, with yearly sampling of gonads to estimate the potential absolute and relative individual fecundity in the Belt Sea (subdivision, SD: 22) and in the Arkona Sea (SD: 24). Additional samples were taken in the Bornholm Sea (SD: 25). The potential absolute fecundity of the total area of investigation is described by the regression models: F_a = 6.89 × TL^3.09 and F_a = ?40 586 + 977.7 × W. As a further result the mean relative fecundity was 978 oocytes per gram body weight. Furthermore, it was shown that there was a difference in development of the potential individual fecundity between the Belt Sea cod and the original eastern Baltic cod stock. In contrast to the stable fecundity of cod in the Belt Sea, the fecundity of the large females in the Bornholm Sea significantly increased from 1993 to 1999. This change is related to a decrease in the dry weight of oocytes in the ovaries. In the same period this cod stock dramatically decreased.     

Sexual maturation of cod (Gadus morhua L.) in the southern Baltic (1990–2006)

Baltic cod, like other species, is susceptible to inter‐annual fluctuations in sexual maturation, depending on the length, age, sex, extent of the habitat area, and stock abundance of the cod population. Maturity is one of the biological indicators used to detect changes in a stock that can be caused by fishing. To address these issues specifically for the eastern Baltic cod stock, long‐term data (1990–2006) from Polish research vessels in the southern Baltic were examined. To date, the ICES has used the same maturity ogives over extended periods and assumed invariant sex ratios for the assessment of eastern Baltic cod. The combined maturity ogives calculated in the present study were markedly lower, particularly for age groups 2–4 (5), in all periods, than those used in the ICES assessment. Moreover, the proportion of females increased with length and age, suggesting that annual verification of the sex ratio is needed. The present study also revealed that the total length (L_50%) and the age (mean age‐at‐maturity; MAM_50%) at which 50% achieved first sexual maturity were higher for females than for males in the study period. The long‐term mean L_50% and MAM_50% for females were 43.9 cm and 4.3 years, respectively, and for males 34.8 cm and 3.4 years. There was also a spatial difference between calculated maturity ogives, with slightly lower L_50% (range: 1.4–8.6 cm) in the Gdańsk Basin than in the Bornholm Basin. The increasing trend in fishing mortality observed in 1993–2004 (ICES data) did not translate into a temporal trend in calculated maturity ogives. However, changes in L_50% and MAM_50% reflected recruitment variations (ICES data). The significance of these findings is discussed in the context of the environment and recruits abundance.     

Assessing seasonal changes in stock composition of Atlantic salmon catches in the Baltic Sea with genetic stock identification

The feasibility of using genetic stock identification to analyse seasonal changes in stock compositions of Atlantic salmon catches in the Baltic Sea was examined. The analysis employed seven variable allozyme loci from most of the potentially contributing stocks (16) from Finland and Sweden. Catch samples were collected from Finnish salmon fisheries in the eastern Bothnian Sea during the 1992 fishing season. Simulation studies were used to evaluate the feasibility of identifying Baltic salmon stocks with allozyme data. Special attention was paid to analysing the wild production of salmon stocks. Clear seasonal differences in stock composition were found. The estimates were compared with smolt production and Carlin-tag data. The proportions of the Neva and Oulujoki river stocks could be estimated as individual stocks, whereas the contributions of the remaining stocks were estimated as four composite stock groups. One of the groups consisted of wild stocks from the rivers Kalixälven and Simojoki. Identification of this group, which could be used as an index of wild production in the catches, requires catch sample sizes >300 salmon if <15% error is required.     

Species and length composition of fish in the south-western Baltic Sea

Length distributions of fish were analysed based on data from the German annual (November/December) bottom trawl surveys in the Mecklenburg Bight and Arkona Basin [ICES Subdivisions (SD) 22 and 24, respectively] of the southwest Baltic Sea, covering the period from 1991 to 2000. The length spectra for all fish species, and those for demersal and pelagic species separately, all showed a consistent pattern of declining logarithmic numbers with increasing length over the fully selected size range. Such consistency allowed the shape of the spectra to be used as a community property that could be tracked in space and time. There was a rapid reduction in the number of specimens in the larger size classes, resulting in steep slopes of the spectra, which may be due to intensive fishing pressure in the Baltic. The species composition of fish communities in the southwest Baltic Sea was stable during the period considered. No consistent species replacement was found, although the clupeid group dropped sharply in the catch during the 1996 survey, which might have been due to sudden environmental changes in that year.     

Ecology and long-term forecasting of sprat (<Emphasis Type="Italic">Sprattus sprattus balticus</Emphasis>) stock in the Baltic Sea: a review

In the brackish Baltic Sea situated in the transition zone between the Atlantic and Euro-Asiatic continental climate systems, the periods of high abundance of sprat of marine boreal origin coincide with a rich freshwater discharge, large water volume inhabitable by fish, rather high winter temperature and low salinity limiting the stock of its main predator—cod (Gadus morhua callarias). In the freshening periods an additional volume of water with acceptable oxygen and temperature conditions for sprat is formed in the Eastern (E) and, especially, the Northwest (NW) regions of the Baltic Sea. This allows extraordinary increase in sprat abundance/biomass. The conditions for sprat deteriorate during the period of active saline water inflows and colder winters, especially in the NW region. Following the decrease in the volume of water acceptable for the wintering of sprat as to temperature and/or oxygen concentration, some part of the stock may be forced to migrate southwards as in the Southwest (SW) region the conditions are the most stable. This may cause an extensive mixing of the stock components of various regions, hindering the differentiation of sprat regional units in the Baltic Sea. As no isolation of regional spawning concentrations has been found, beginning with 1989 the Baltic sprat has been assessed and managed as one stock unit, despite spatial differences in its morphological characters, growth rate etc. The alternation of periods of different regime-forming conditions (freshening or oceanization) is probably triggered by climate changes. This regularity has been exploited for the composition of long-term forecasts of qualitative changes in fish (incl. sprat) stocks in the Baltic Sea.     

The stock structure and migrations of plaice populations on the west coast of England and Wales

Plaice Pleuronectes platessa populations on the west coast of England and Wales are currently managed as two stocks: in ICES Division VIIa (Irish Sea, Cardigan Bay and St George's Channel), and ICES Divisions VIIf&g (Bristol Channel and Celtic Sea). A total of 13,784 plaice were fitted with Petersen tags and released in these areas during 1979–1980 and 1993–1996. Analysis of the 2788 recaptures received by June 2000 confirmed known spawning and feeding grounds in the region. It showed also that plaice >25 cm L _T tended to undertake extensive spatial movements. At this size, female plaice were likely to be mature or maturing for the first time, whilst males were probably mature. Tag recaptures indicated resident sub-stocks of plaice in the north-east Irish Sea, the south-east Irish Sea, Cardigan Bay and Bristol Channel, a contingent of plaice in all areas that undertook permanent dispersal to other areas, and a contingent which originated in the south-east Irish Sea and migrated to spawn in the Bristol Channel. Plaice originating in the Bristol Channel rarely moved north into ICES Vila. A general hypothesis of plaice population structure in the region is presented and discussed in relation to stock assessment.     

Impact of hypoxia on consumption of Baltic cod in a multispecies stock assessment context

The Baltic Sea is characterised by a heterogeneous oceanographic environment. The deep water layers forming the habitat of Baltic cod (Gadus morhua callarias L.) are subjected to frequently occurring pronounced anoxic conditions. Adverse oxygen conditions result in physiological stress for organisms living under these conditions. For cod e.g. a direct relationship between oxygen availability and food intake with a decreasing ingestion rate at hypoxia could be revealed. In the present study, the effects of oxygen deficiency on consumption rates were investigated and how these translate to stock size estimates in multi‐species models. Based on results from laboratory experiments, a model was fitted to evacuation rates at different oxygen levels and integrated into the existing consumption model for Baltic cod. Individual mean oxygen corrected consumption rates were 0.1–10.9% lower than the uncorrected ones. At the currently low predator stock size, however, the effect of oxygen‐reduced consumption on the total amount of eaten prey biomass and thus predation mortalities was only marginal. But should successful management lead to higher cod stock sizes in the future, then total predation mortalities will greatly increase and thus improved precision of these estimates would be valuable for the assessment of prey stocks.     

Stock‐specific variation of trophic position,diet and environmental stress markers in Atlantic salmon Salmo salar during feeding migrations in the Baltic Sea

This study investigated stock‐specific variation in selected ecophysiological variables during the feeding migrations of Atlantic salmon Salmo salar in the Baltic Sea. Oxidative stress biomarkers and EROD (ethoxyresorufin‐O‐deethylase, Cyp1A enzyme) activity were used as indicators of possible environmental stress and stable isotopes as determinants of diet and trophic position. Latvian S. salar stocks Daugava and Gauja had distinct stable‐isotope signatures compared to the other stocks, indicating differences in migration patterns, residency or arrival times, or dietary specialization among stocks. Salmo salar originating from Daugava and Gauja also had lower catalase enzyme activity than the other stocks. Post‐smolts originating from rivers of the Gulf of Finland had elevated EROD activities compared to fish of the same age from Bothnian Bay rivers, which could indicate exposure to organochlorine pollutants. No other stock‐specific differences in oxidative stress biomarkers were found. The study demonstrates how genetic, oxidative stress biomarker, EROD and stable‐isotope data may be combined to study trophic position, prey prevalence and environmental stress of mixed S. salar stocks foraging in the sea.     

The estimation of a mackerel (Scomber scombrus L.) spawning stock size by plankton survey

The International Council for the Exploration of the Sea (ICES)Mackerel Working Group recognises two mackerel stocks in Europeanwaters, the North Sea stock and the Western stock. From Marchto July 1977 an Anglo-French plankton survey was carried outin the shelf waters of the Bay of Biscay, Celtic Sea and westof Ireland to estimate the size of the Western stock. Sampleswere collected with a 76 cm Lowestoft pattern high-speed planktonsampler fished to a maximum depth of 100 m. The number of eggsm^–2 at each of five development stages were raised todaily production estimates with development rate coefficientsmeasured with live material collected in March. The mean numberof stage I eggs m^–2 d^–1 was raised to a total dailyproduction estimate for each month by the area within the minimumcontour level, and a production curve was drawn for the spawningseason. The total egg production was estimated at 1.98 x 10¹⁵eggs, + 30%, – 20%. With a mean fecundity of 361 000 anda sex ratio of 0.64 males per female this is equivalent to 8995 million fish.     

Effect of spatial differences in growth on distribution of seasonally co‐occurring herring Clupea harengus stocks

The mechanisms most likely to determine the distribution of the two major herring Clupea harengus stocks in their common early summer feeding ground in the eastern North Sea, Skagerrak and Kattegat were investigated through analysis of acoustic survey data from six consecutive years. No change was detected in biomass of North Sea autumn spawning C. harengus (NSAS) over time, whereas the biomass of western Baltic spring spawning C. harengus (WBSS) declined severely. Analyses of centre of abundance by stock showed no change in NSAS distribution, whereas the WBSS changed to a more western distribution over time. Contrary to previous perception of the juvenile migration, NSAS were found to leave the study area at the age between 1 and 2 years and WBSS 1 year olds were encountered in the Skagerrak. The estimated parameters of von Bertalanffy growth equations showed marked differences between areas with fish in the eastern part of the area having the lowest size at age at all ages. Further, their growth conditions appeared to deteriorate progressively over the period studied. Both NSAS and WBSS showed the highest condition in the North Sea and Skagerrak while condition was substantially lower in age Kattegat. The westward movement of spring spawners over time suggests that growth rate and possibly density of conspecifics influence the migration pattern and distribution of C. harengus in the area. In contrast, there was no evidence to suggest that distribution was constant over time within stocks or that distribution reflected size‐dependent limitations on migration distance.     

An outbreak of type C botulism in herring gulls (Larus argentatus) in southeastern Sweden

From 2000 to 2004, over 10,000 seabirds, primarily Herring Gulls (Larus argentatus), died from an undetermined cause in the Blekinge archipelago in southeastern Sweden. In June 2004, 24 affected Herring Gulls were examined clinically, killed humanely, and 23 were examined by necropsy. Seven and 10 unaffected Herring Gulls collected from a local landfill site and from Iceland, respectively, served as controls. All affected birds showed similar neurologic signs, ranging from mild incoordination and weakness to severe flaccid paralysis of legs and wings, but generally were alert and responsive. All affected gulls were in normal nutritional condition, but were dehydrated and had empty stomachs. No gross or microscopic lesions, and no bacterial or viral pathogens were identified. Type C botulinum toxin was detected in the sera of 11 of 16 (69%) affected gulls by mouse inoculation. Type C botulism was the proximate cause of disease in 2004. Sera from 31% of birds tested from outbreaks in 2000 to 2003 also had detectable type C botulinum toxin by mouse inoculation. No large-scale botulism outbreak has been documented previously in this area. The source of toxin, initiating conditions, and thus, the ultimate cause of this outbreak are not known. This epidemic might signal environmental change in the Baltic Sea.